taxonID	type	description	language	source
2D248795FFBFE14AFF64F92101F5F8FB.taxon	description	(Figures 1 – 8, 10 B)	en	Keetapithchayakul, Tosaphol Saetung, Kim, Junggon, Phan, Quoc Toan, Digma, Jonathan Rodrin (2026): Reexamination of Heteronaias heterodoxa (Selys, 1878) (Odonata: Corduliidae), with larval redescription, comparative notes on Asian Corduliidae larvae, and biogeographic significance. Zootaxa 5777 (3): 471-488, DOI: 10.11646/zootaxa.5777.3.4, URL: https://doi.org/10.11646/zootaxa.5777.3.4
2D248795FFBFE14AFF64F92101F5F8FB.taxon	materials_examined	Material examined. PHILIPPINES. 4 adults: 1 ♂, 1 ♀ Calanasan stream (18 ° 24 ' 16.8 " N, 121 ° 03 ' 18.2 " E, elevation 645 m a. s. l.) Calanasan, Apayao, 12 / X / 2025, J. M. Garces leg.; 1 ♂, upstream Binangawan fall (09 ° 09 ' 25 '' N, 124 ° 43 ' 57 '' E, elevation 900 m a. s. l.), Sagay, Mt. Mambajao, Camiguin, 9 / XII / 2018, H. Freitag leg.; 1 ♀, lower reach of Taugad River (12 ° 37 ' 25 " N, 124 ° 43 ' 57 " E, elevation 140 m a. s. l.), Brgy. San Vicente, Roxas, Oriental Mindoro, 28 / XI / 2017, J. M. Garces leg. 1 exuviae: 1 ♀, (as last stadium larvae, reared), Calanasan stream (18 ° 24 ' 16.8 " N, 121 ° 03 ' 18.2 " E, elevation 645 m a. s. l.), Calanasan, Apayao, 12 / X / 2025, J. M. Garces leg. 10 last stadium larvae: 1 ♂, 1 ♀, Sitio Setic Barangay, Ibona, Ibona Town (15 ° 15 ' 08.3 " N, 121 ° 23 ' 30.2 " E, elevation 44 m a. s. l.), Dingalan, Aurora, 16 / III / 2025, J. Rocha leg.; 1 ♂, 1 ♀ Calanasan stream (18 ° 24 ' 16.8 " N, 121 ° 03 ' 18.2 " E, elevation 645 m a. s. l.), Calanasan, Apayao, 12 / X / 2025, T. S. Keetapithchayakul leg.; 1 ♂, Mimpis River (18 ° 24 ' 16.8 " N, 121 ° 03 ' 18.2 " E, elevation 645 m a. s. l.), Bukal, Majayjay, Laguna, Luzon, 28 / VI / 2025, BIO-PHIL exped.; 1 ♂, upstream Binangawan fall (09 ° 09 ' 25 '' N, 124 ° 43 ' 57 '' E, elevation 900 m a. s. l.), Sagay, Mt. Mambajao, Camiguin, 9 / XII / 2018, H. Freitag leg.; 1 ♂, 1 ♀, lower reach of Taugad River (12 ° 37.30 ' N, 121 ° 22.97 ' E, elevation 140 m a. s. l.), Brgy. San Vicente, Roxas, Oriental Mindoro, 28 / XI / 2019, H. Freitag leg.; 1 ♂, 1 ♀, Tagbak (8 ° 00 ' 26 " N, 124 ° 41 ' 24 " E, elevation 1210 m a. s. l.), Talakag, Bukidnon, Mindanao, 16 / VIII / 2023, H. Freitag leg. Notes on adult morphology. The male and female adult specimens of H. heterodoxa matched well with the species diagnoses of Needham & Gyger (1937) and Asahina (1980), except where indicated. Male: body shining metallic green (Fig. 1 A – B); vesica spermalis (Fig. 1 C – D) segment 4: lateral lobes large, elongate, projecting ventrally and slightly curved posteriorly, forming a prominent chitinous blade-like expansion; medial lobe bilobed, distinctly broadened and with two rounded lateral swellings separated medially; medial process divided into two distinct portions, the upper portion broad and chitinous-like at the base, with lateral margins slightly expanded, gradually narrowing distally, each side of distal portion with a backward coiled laterodistal flagella, the lower portion stout basally, with distal end produced into a dorsally curved filament. Anal appendages (Fig. 1 E – G) as long as 9 and 10 together: cerci elongate, slender, tapering to blunt tips, diverging distally in dorsal view, slightly curved in lateral view, dense SS along margins, ventral view with distinct protuberance; epiproct distinctly shorter than cerci (about two-thirds their length), elongate and slender, basally broad and gradually tapering, in lateral view slightly convex on ventral margin, and with distal end truncate, bearing a pair of small tubercles; margins with sparse SS. Female: similar to male unless otherwise stated (Fig. 2 A – B). Hind wing without anal triangle and anal angle. Posterior end of abdomen broadly expanded laterally from S 9 to S 10, forming broad sub-sagittal lobes that extend ventrally the anal appendages (Fig. 2 C – F). Cerci black, elongate, slender, apices acute, margins densely covered with long SS; in dorsal view diverging slightly, in lateral view ventral margin weakly curved dorsally. Vulvar lamina U-shaped, with lateral lobes triangular and acute in shape.	en	Keetapithchayakul, Tosaphol Saetung, Kim, Junggon, Phan, Quoc Toan, Digma, Jonathan Rodrin (2026): Reexamination of Heteronaias heterodoxa (Selys, 1878) (Odonata: Corduliidae), with larval redescription, comparative notes on Asian Corduliidae larvae, and biogeographic significance. Zootaxa 5777 (3): 471-488, DOI: 10.11646/zootaxa.5777.3.4, URL: https://doi.org/10.11646/zootaxa.5777.3.4
2D248795FFBFE14AFF64F92101F5F8FB.taxon	description	Redescription of larva. Medium size, body robust, moderately flattened, anal pyramid blunt, and coloration pale yellowish brown to reddish brown (Fig. 3). Head. Wider than long, about 1.44 times as wide as long (Figs 3, 4 B). Labrum broad and flattened ventrally, with the ventral margin concave, scattered long SS on anterior and dense SS on middle to distal part. Clypeus (Fig. 4 B) broad and flattened ventrally, with the ventral margin shallowly concave, mostly glabrous. Antefrons slightly convex, with scattered small tubercles, flattened. Postfrons wide and slightly convex, extending laterally to the inner margins of compound eyes, anterior margin gently curved, merging indistinctly with antefrons, posterior margin extending laterally to the inner margins of compound eyes, distinctly bordered by a broad, dark band along postfrontal suture, interrupted medially by three ocelli and a narrow zone of fine longitudinal striations which separate the left and right dark patches. Antennae (Fig. 4 A – D) 7 - segmented; A 3 the longest segment, relative lengths of antennomeres: 0.14: 0.15: 0.3: 0.13: 0.11: 0.10: 0.07. A 1 and A 2 short, barrel-shaped, A 3 elongate and slender; A 4 – 7 gradually tapering distally. All segments (A 1 – 7) covered with scattered SS and small tubercles. Compound eyes (Figs 3 A, 4 B, 4 D) moderately developed, protruding latero-dorsally. Vertex broad and moderately convex, posterior margin slightly emarginate at midline; occipital lobe (Fig. 4 B) lateral margins almost straight, strongly convergent posteriorly, extending posteriorly and ventrally beneath the compound eye, with a distinct crescent-shaped pale dark patch often present along the ventral margin. Genae (Fig. 4 D). Genae (Fig. 4 C, 4 E) smooth, with a distinct row of stout PS along lateroventral margin below the compound eyes, with tufts of long SS and a distinct lateral row of stout PS near maxillary base. Labium (Figs 3 A, 5 A – B) spoon-shaped; prementum-postmentum articulation reaching anterior part of procoxae. Prementum (Fig. 5 A – B) 1.8 × wider than long, pentagonal shaped, latero-basal margins slightly concave; in dorsal view with dorsolateral rows of patches of SS and intermingling row of 8 – 10 SPS among them, with laterodistal end with 2 – 4 stout SPS, and with anterodorsal area with a dense patch of 20 – 30 SS near labial palp articulation; premental setae with 9 – 10 long and 2 – 4 short setae (9 + 4, 10 + 2) (sometimes 8 long setae) to each side of midline; ventral surface glabrous, with a short central sulcus; distal margin of prementum (Fig. 5 C) well developed transversally, but poorly anteriorly, margin only slightly convex, with a row of 13 raptorial setae, distal marginal end with 11 – 12 shallow crenulations, each crenulation finely serrated, with the 2 – 6 SPS on each lobe. Labial palp (Fig. 5 D – F) strongly developed, covering anterior part of head as a mask, covering clypeus, the apical margin with six large, finger-like round-tipped teeth of different size, all of them finely serrated on margins, distal ends with raptorial setae as follows from dorsum to venter: L 2: 2: 2: 3: 3: 12, R 1: 2: 2: 3: 3: 12; dorsal margins with a row of small SPS; ventral margins with small inconspicuous serrations and 4 – 5 long SPS; with 10 long palpal setae, one basal setella and a group of 7 – 8 small spiniform setae close to articulation; movable hook slender, sharply pointed, smooth. Mandibles (Fig. 6 A – H) with formula: L 1234 0 a b, with incisors 1> 2 <3 = 4 / R 1234 y a b, with incisors 1> 2 <3 <4, asymmetrical, robust with well-developed long teeth; outer margin of mandible with median portion glabrous and surrounding three patches of dense, long SS. Maxilla (Fig. 6 I – K) galeolacinia with seven teeth, four dorsal teeth of approximately the same size, apical tooth largest, three ventral teeth of different size, the basal tooth smallest. Thorax. Prothorax (Figs 3, 7 A – B) narrower than head, covered densely by minute tubercles; notum almost square in shape, with anterior ridge biconvex, a row of long SS on the central part, lateral margins rounded, posterior ridge biconvex and with a row of long SS along its inner margin between middle groove, posterior margin forming a subtle ridge across posterior border and with a row of long SS; pleura (Fig. 7 A – B) with a strongly developed episternal apophysis, digitiform, paddle-like in anterior or posterior view, broad at base, tapering distally with rounded tip, gently curved anteriorly; inferior margin of epimeron gently convex. Pterothorax (Fig. 3) broader than pronotum, dorsal area of mesopleura shield-like, with a row of long SS along lateral ridge, with a pair of large, transverse spiracles on most anterior part. Wing sheaths parallel; anterior and posterior wing sheaths reaching respectively posterior margin of S 5 and basal half of S 6. Legs (Fig. 7 C – E) coxa robust; femora thick, strongly depressed, covered by a row of long SS on outer margin; tibia depressed, more slender than femora, slightly curved, with a longitudinal line on dorsal view, with four distinct longitudinal rows of small SPS, arranged in parallel, extending to distal end on ventral view; tibial comb (Fig. 7 F – H) densely covered by short SS and SPS on distal end; tarsal formula: 3 – 3 – 3; with scattered 2 rows of fine, robust, SPS on ventral surface of each segment, tarsomere II and III with respectively a pair of stout spines and distinct SS on distal end; pretarsal claws strongly curved, with developed empodium. Abdomen. Ovate, widest at S 6, abruptly narrowed posteriorly on S 10, strongly convex dorsally, and flat ventrally. Abdominal terga consisting of tufts of SS on mesolateral area of S 2 – 6 and posterolateral area of S 6 – 7; medial posterior margin of S 3 – 6 with a dense row of small SS; S 6 – 10 with middorsal protuberances; posterior margin of S 5 – 7 with a row of long SS, S 8 – 9 with short SS along posterior margin; lateral margins of S 8 – 9 ending in a slightly convergent, blunt spines, S 8 spines small, S 9 spines very large (Fig. 8). S 10 short and sunk into S 9, Sterna S 1 – 10 (Fig. 8 A) covered abundantly with small SPS; S 10 hardly visible in ventral view posterior margins of sterna S 9 with a row of dense and long SS; the secondary genitalia on the sterna of S 2 – 3 visible through the transparent cuticle. Antepleurites present and rather well developed on S 3 – 4. Female gonapophyses pads vestigial, bilobed, lobes conical with apices rounded, the space in between equal to the basal width of each lobe on S 9. Anal pyramid (Fig. 8 B – F), blunt, pyramidal shape; epiproct as long as paraprocts, with scattered distinct SPS on ventral view, with a low dorsomedial tubercle and small convex distal margin in male; paraprocts with medial carina bearing dense SS on inner side; cerci conical, 0.76 as long as epiproct length and 0.51 in female. Larval variation (n = 10). Relative lengths of antennomeres following (0.05 - 0.14): (0.08 - 0.15): (0.19 - 0.3): (0.06 - 0.13): (0.08 - 0.11): (0.05 - 0.1): (0.04 - 0.07). Premental setae vary from 9 + 9, 10 + 9, 9 + 10, 10 + 10, 11 + 10, 11 + 11, 11 + 12, 12 + 13, and 13 + 11. Palpal setae vary from 8 – 11. Apical margin of labial palp forming 5 – 7 lobe, each lobe varied raptorial setae 1 – 3 and the last lobe (ventre one) vary 10 – 14.	en	Keetapithchayakul, Tosaphol Saetung, Kim, Junggon, Phan, Quoc Toan, Digma, Jonathan Rodrin (2026): Reexamination of Heteronaias heterodoxa (Selys, 1878) (Odonata: Corduliidae), with larval redescription, comparative notes on Asian Corduliidae larvae, and biogeographic significance. Zootaxa 5777 (3): 471-488, DOI: 10.11646/zootaxa.5777.3.4, URL: https://doi.org/10.11646/zootaxa.5777.3.4
2D248795FFBFE14AFF64F92101F5F8FB.taxon	distribution	Distribution. Heteronaias heterodoxa occurs across the Philippine archipelago, with numerous records documented from Luzon, the Visayas, and Mindanao (Fig. 9). Although Asahina (1980) reported a single specimen from Palawan, some doubt exists over this record. Habitat and biology. The larvae of H. heterodoxa inhabit shaded forest streams, from lowland to mid-elevation tropical evergreen forests (Fig. 10 A). The streams are characterized by clear, shallow water running over a substrate composed primarily of boulders and large stones (> 25 cm) (40 %), pebbles (30 %), leaf litter and organic debris (20 %), and sand / silt (10 %). Larvae were discovered on the undersides of rocks and stones measuring approximately 15 – 35 cm in diameter, in slow to moderately flowing water. Exuviae were observed attached to inclined or vertical surfaces of large rocks above the waterline (Fig. 10 B), suggesting that final instar larvae actively crawl out of the water, anchoring themselves to stable surfaces for adult emergence.	en	Keetapithchayakul, Tosaphol Saetung, Kim, Junggon, Phan, Quoc Toan, Digma, Jonathan Rodrin (2026): Reexamination of Heteronaias heterodoxa (Selys, 1878) (Odonata: Corduliidae), with larval redescription, comparative notes on Asian Corduliidae larvae, and biogeographic significance. Zootaxa 5777 (3): 471-488, DOI: 10.11646/zootaxa.5777.3.4, URL: https://doi.org/10.11646/zootaxa.5777.3.4
